The Desert Stink Beetles in the tribe Amphidorini LeConte (genus Eleodes Eschscholtz and relatives) represent a diverse lineage of flightless beetles well-adapted to the arid regions of western North America. Specimen data from natural history collections has been combined with molecular phylogenetics to explore the evolutionary and biogeographic history of the tribe. These data are then used to explore how deserts, and particularly sand-dune systems, should divided into biogoegraphic regions in the desert southwest.
Biogeography and Evolution of the Desert Stink Beetles
1. Biogeography and Evolution of the
Desert Stink Beetles
(Coleoptera: Tenebrionidae: Amphidorini)
M. Andrew Johnston
2.
3.
4. AmphidoriniLeConte, 1862*
• New World tribe
• 7 Genera
– 17 subgenera of Eleodes Eschscholtz
• 256 valid species/subspecies
– 213 in Eleodes, with 163 additional
synonyms
= Eleodini Blaisdell, 1909
Eleodes spinipes macrura Champion, 1892
5. AmphidoriniLeConte, 1862
• Last comprehensive revision was
Blaisdell, 1909
• Treated 115 species/subspecies
from the U.S. and Canada
Eleodes snowi Blaisdell, 1909
6. AmphidoriniLeConte, 1862
• All species are flightless
• Known for defensive “head-
standing” posture
Eleodes hispilabris (Say, 1824)
7. Several sympatric species found in Arizona
E. madrensis Johnston
E. subnitens LeConte E. longicollis LeConteE. carbonaria Say
E. anthracina Blaisdell
Eleodes Eschscholtz, 1829
8. Müllerian mimicry
Collected at same time
(Heber, AZ 2013)
Smith et al. 2014, fig. 2
Morphological
convergence
• Sculpturing
• Dorsal silhouette
9. Batesian Mimicry
• 2 are Stenomorpha
• Lack defensive
glands
• Mimic
headstanding
behavior
• ~180 my divergence
Collected at same time
(Heber, AZ 2013)
Smith et al. 2014, fig. 2
10. Biodiversity data and darkling beetles
Eleodes Eschscholtz, 1829
213 Valid species-group taxa
412 Available species-group names
22. Failure across (at least) Coleoptera
iPhylo Blog Post
iPhylo.blogspot.com
~85,000 Australian Coleoptera records
1/3 names are changed, affecting 1/5 records
23. Graduate students should be empowered
to use and curate biodiversity data
Backbone taxonomies and early-career systematists
24. Graduate students should be empowered
to use and curate biodiversity data
Backbone taxonomies and early-career systematists
Current backbone taxonomies alienate systematists
25. Graduate students should be empowered
to use and curate biodiversity data
Backbone taxonomies and early-career systematists
Need to develop new, community-empowering designs
Current backbone taxonomies alienate systematists
37. Checklist generated from digitized records
Currently digitized data are useful, though not complete
38. Checklist generated from digitized records
Currently digitized data are useful, though not complete
Faunal and revisionary studies should
explicitly reference available records
39. Checklist generated from digitized records
Currently digitized data are useful, though not complete
Faunal and revisionary studies should
explicitly reference available records
Taxonomic expertise is still very much required
43. Current Generic Classification
Phylogenetic analysis of Amphidorini
South American Nycterinus Eschscholtz
does not belong in Amphidorini
‘True Amphidorini’ are only found in
North America
Nycterinus rugiceps Curtis, 1845
44. Phylogenetic analysis of Amphidorini
Paraproct
Valvifer
(Coxite I)
Gonostyle
Coxite IV
Coxite III
Coxite II
Nycterinus rugiceps Curtis, 1845 Eleodes armata LeConte, 1851
47. Consensus tree of 500 ML replicates
7-Gene Phylogeny Backbone
is Poorly Supported
7-way Polytomy 5-way Polytomy
Phylogenetic analysis of Amphidorini
48. Supports our 7-gene
topology
Currently analyzing
29 transcriptomes
from all major
lineages
Preliminary 343-loci analysis
Phylogenetic analysis of Amphidorini
50. Comparative Morphology
Many molecular clades are
finding morphological support
Female ovipositor characters
can diagnose subgenera
Female Ovipositors of 3 different subgenera,
A,B: E. tribula, C: E. barbata, D: E. hirsuta
56. CORRELATION*
*Landrum, L.R. and D. Lafferty. 2015. Taxon 64(5) 998-1016
http://dx.doi.org/10.12705/645.9
Computes how frequently each species is found
near each other given species
57. CORRELATION*
*Landrum, L.R. and D. Lafferty. 2015. Taxon 64(5) 998-1016
http://dx.doi.org/10.12705/645.9
Clusters species into co-occurring groups
58. CORRELATION
*Brown, D.E. (ed.) 1994. Biotic communities: Southwestern United States and northwestern
Mexico. Salt Lake City: University of Utah Press.
Arizona Flora Test Case:
• Examined 81 ‘high
profile’ species
• Recovered groupings
highly congruent with
published biotic
communities*
59. CORRELATION
Analyze input set of
lat/long coordinates:
• Correlate an animal
species with SEINet
plants
• Place that animal in a
plant-driven biotic
community
60. Using CORRELATION, we can
ask questions about ecological
communities
And answer them based on
specimen occurrence data
61. Where can we find a certain species?
“You can find it in
oak-juniper forests in
southern Arizona”
Eleodes madrensis Johnston, 2015
62. Where can we find a certain species?
Eleodes madrensis Johnston, 2015
5 highest correlated plant
species
“You can find it in
oak-juniper forests in
southern Arizona”
65. Differentiate habitats of similar species
Eleodes arcuata Casey, 1884Eleodes debilis Horn, 1870
Only share a single species of top 10
66. Differentiate habitats of similar species
Eleodes arcuata Casey, 1884
Eleodes arcuata is
correlated with oak-
juniper mid-elevation
transition forest
67. Differentiate habitats of similar species
Eleodes debilis Horn, 1870
Eleodes debilis is
correlated with riparian
broadleaf plants from a
similar elevation
68. CORRELATION transforms our approach to
understanding distribution
Ecological Community
“Authoritative observation”
Repeatable, specimen-based
correlation
69. Ecological Community
“Authoritative observation”
Repeatable, specimen-based
correlation
Spatial Distribution
Geographic polygons
Specimen-driven, co-occurring
species communities
CORRELATION transforms our approach to
understanding distribution
70. Can we infer ancestor-descendant
plant-community shifts?
71. Historical plant-community inference
• 21 Eleodes species from Arizona were selected
– Records with high geographic uncertainty or missing
locality data were removed
– Records were sorted to taxa by ‘ScientificName’ based on
nomenclatural synonymies
72. Historical plant-community inference
• 21 Eleodes species from Arizona were selected
• Associated beetles with plant communities using
CORRELATION
– Beetle species associated with 6 different plant
communities
– 18 species were correlated to a single plant community, 3
were equally correlated to 2 communities
73. Historical plant-community inference
• 21 Eleodes species from Arizona were selected
• Associated beetles with plant communities using
CORRELATION
• ‘Biogeographic’ reconstruction in BioGeoBEARS
– Inferred historical plant-community lineage associations
– Assumed current plant communities existed throughout
Eleodes evolutionary history
75. Historical plant-community inference
Two independent
radiations into the
Sonoran Desert
community
Phylogeny based on a 7-gene unpublished
dataset, Smith et al. in prep
76. Historical plant-community inference
‘Melaneleodes Clade’
ancestors associated
with Pine-Juniper
community of the
Colorado Plateau
Phylogeny based on a 7-gene unpublished
dataset, Smith et al. in prep
77. Historical plant-community inference
Ancestral lineage
associated with higher
elevation oak-juniper
and ponderosa pine
communities
Phylogeny based on a 7-gene unpublished
dataset, Smith et al. in prep
78. • Novel analysis for inferring historical habitat
transitions
• Entirely based on vouchered specimen data
• Analyses are reproducible and easily re-done when
new data are made available
Historical plant-community inference
79. Trogloderus LeConte, 1879
• Endemic to sand dunes
and sandy habitats in
western United States
• Currently comprised of 1
species with 4 subspecies
Trogloderus ‘major’ n.sp.
80. Last revised by La Rivers in
1946
• Invoked orthogenetic evolution
81. Last revised by La Rivers in
1946
• Invoked orthogenetic evolution
• Hypothesized Trogloderus to be ancient
82. Last revised by La Rivers in
1946
• Invoked orthogenetic evolution
• Hypothesized Trogloderus to be ancient
• Hypothesized origins from the Great Basin
83. Phylogenetic revision of Trogloderus
36 OTUs, 6 loci
analyzed in BEAST
• 10 total species
• 4 described, 6 new
• Trogloderus perhaps
~5.5 my old
Trogloderus root
87. Van Dam and Matzke 2016
What about classifying sand dune regions?
88. Trogloderus as a biogeographic model
• Moderate habitat specificity
• Sandy, well-drained soils
• Apparently wide temperature tolerance
• Low to moderate dispersal
• Flightless
• Not restricted to single localities
• Species show strong geographic signal
90. Great Basin
T. costatus LeConte
Great Basin
T. nevadus La Rivers
Prehistoric Lake Subregions
• Lake Lahontan (W)
• Lake Bonneville (E)
91. Great Basin – Mohave Transition
T. ‘kandai’ n. sp.
T. ‘arcanus’ n. sp.
T. ‘major’ n. sp.
92. Great Basin – Mohave Transition
T. ‘kandai’ n. sp.
T. ‘arcanus’ n. sp.
T. ‘major’ n. sp.
“Great Basin”
“Mohave Desert”
93. Lahontan Trough
Proposed by Reveal in 1979
• Low-elevation desert
corridor
• Geologically distinct from
Lake Lahontan
94. Lahontan Trough
Eusattus muricatus LeConte
• Genetically distinct population1
• First to recognize Lahontan Trough
as a genetic barrier
1Britten & Rust 1996. Conservation Biology
95. Lahontan Trough
Eusattus muricatus LeConte
• Genetically distinct population1
• First to recognize Lahontan Trough
as a genetic barrier
E. muricatus species group
• Exhibit speciation consistent with
the Lahontan Trough2
1Britten & Rust 1996. Conservation Biology
2Doyen 1984. Occ. Papers of Cal. Acad. Sci.
96. Owens Lake Basin
T. ‘kandai’ n. sp.
T. ‘arcanus’ n. sp.
T. ‘major’ n. sp.
“Great Basin”
“Mohave Desert”
• Southern extent of ‘Great
Basin’ flora
• Not connected to Lahontan
Trough
• T. ‘kandai’ is the first
reported endemic sand
obligate
100. Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
101. Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
102. Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
– Likely relatively young, roughly 5–6my old
103. Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
– Likely relatively young, roughly 5–6my old
• Hypothesized origins from the Great Basin
104. Revisiting La Rivers 70 years later
• Invoked orthogenetic evolution
– Well-adapted to cryptic living in sand
• Hypothesized Trogloderus to be ancient
– Likely relatively young, roughly 5–6my old
• Hypothesized origins from the Great Basin
– Probably true; Great Basin younger than he knew
105. Trogloderus Biogeography Insights
• Intermountain Region has produced complex
speciation during the recent past
• Transition zones between broader ecoregions
could be their own areas of endemism and
speciation
106. Collaborators
Nico Franz
Sal Anzaldo
Andrew Jansen
Brian Reiley
Bill Warner
Fred Skillman
Kojun Kanda
Aaron Smith
Sangmi Lee
Liz Makings
Chris Wirth
Ryan Lume
Travis Hitchner
Brennan Hayes
David Flemming
Monica Chacon
CMN Visiting Scientist Fellowship
CanaColl
Foundation
NSF ARTS DEB-1258154
Funding
Thank you!
Editor's Notes
Doyen and Somerby 1974
Perhaps the most majestic group of all animals
Grammatical gender – nomenclaturally complex group
Header – taxonomic historiy/issues etc
Issues I, then II next slide
Mention literal text copy
Harvested and available via GBIF
Verbatim records are available, but then you need taxonomic expertise to decode
Not issues with data quality, but with backbone taxonomic filtering
EARLY CAREER SYSTEMATISTS - FRUSTRATION
Building global initiatives / infrastructure – should be used by and encourage participation from E.C. researchers